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The Hole of Burrowing Sponges Ljl Bioerosion· Ocoologill (Berl.) 19, 203-21(j (1975) (C by Springer.Verlllg 1975 The Hole of Burrowing Sponges lJl Bioerosion· Klaus Ri,ihler Department of 'nvertebrato Zoology. National Museum of Naruml History, Smithwnian IMtitution. Washington, D.C. 205G0 Heceived February 10, 1975 Summary. Among the lurge number of limestone.eroding organisms. sponge8, mninl;)' of the ramily Clionidac are of special intctellt be<:llulle of thcir efficient meHnll of subl!tmtum penetration by cellular ctching and bccUUIIC they relellsll clmrlleh::rilltiCHlly shaped calcium carbonate chipll which call be delccted in tho mud·size fraetion or lIIany 8Cdimentl'l. Tdenti· finble tTlU:'C f08lliia nud lIt..dimenhl lire of great ecological Ilnd pHleoecologicll1 significllllce. As new data on the cXCllvllliug mechanism 1111\'8 become available, the qUClltiollll of burrowing mlt:tlllnd llt..diment production 11Il\'c gained importance. E"tMlpolation from short­ term cxperiments (undcr 6 months) on substrate ill\'llsion are inoolll::lusive be<:aUJIll or high initilll peuctmtion mtCll fClIulling from nll.'chauical stimulation llnd lack of competition. New experimcnts show thllt the rate CUT\'C f1attclls after (j monthll lind thllt optimum long, tcrm erosion of Onco, dOCll not exCt.'(..,J 700 mg m-2 ycar-l (Cliolla lampa llnd C. uprica). Subs~nlt6 limil.fttioll.!llUld competition will furthcr reduce this rate. By monitoring the production of CnCO, chips by CIiOllU lampo.. it "'all polIIIible to link tlCtivit;r patterns to certain eUVirollnlClllt,1 factors. Medmnical stimuli, high light intcnllity. strong currents lind, possibly, low tcmperature llcem 10 11Ct.'Clerntc the burrowing procell8. Sponge.gencrated chipIJ can IIlliko up Ol'er 40% or oond mud when deposited in tho current shadow of the reef framework. Using tmnsoot counts and sponge 1lrea,·biomll/lS oolll'cl1Iion fnctol1l. the moon abundllnce of burroWing spcmgCEl on the Bermudll pl,ltform could be cn1eulnted. On suitablo hllrd bottom substmtCll it averagC6 W g dry weight per m". From thia \'lIlue the burrowing potcntilll of 2~;(j spongCEl can be catimatcd lUI g CllCOI per m' ilubstrnt:c per ycnr, Since 97-98% of the eroded limestone remnins in particulate form, the contribution of fine scdimcnlJl can nmount to 250 g m-' yearl, Attention is called to the fact that erosion mtes by burrowel1l can not dii'QCtly be oom­ parod with thOBO of borers or scrapers. Tho former lire intermittent and their activitiC6 lire affooted by environmentalnnd biological inlcractions, while activities of the Intter lire rather constant lind guided by the noed for food. Inlroduction Sponges of tbe family Clionidac have long boon known for the ecological impact they cause by riddling limcstome objcct-s which thcy inhabit in most marine environments. Damagc t-o oyster cultures by clionids has been publicized since the early part of the Ninetccnth Century (references in: Clapp and Kcnk, 1963) and is still stimulating Slllyeys and experimental ftJ!lCarch (Hopkins, 1956; Hartman, 1958; Hoese and Durant, 1968). • Thi~ work Wllij supporled h~' lhe SlIlith~onilln RCliClirch Foundation. by the Smithsonian Environmental $cicnCCll Program and by n Sydney I~. Wright Fellowship. lit the Bermuda Biological Station, Contrihution No. (j11J, .Bernllldll Biological Station. Contribution No. 10. Investigations of Milrine Shallow Water EOOllystenlS Project, Smith80nilln I.alltitution. '''' K. Riltzlcr Ecological requiremcnt.s of CliO/la, particularly salinity and temperature tolerances, competition for food nnd substrata llnd dept.h dislribul·ion were studied llnd discussed by Volz (1939), Hart.man (1957, 1958), Driscoll (1967), Bromlcy and Tcndal (1973) and Pang (1973). Possible usc of these dnta for paleoecological analyses nnd for life Ilnd post mort-em histories of shell bearing invertebrate fossils was proposed by Lawrence (1969) and Bromley (1970). The burrowing mechaniSlll used by sponges, an old controversy between advocntes of chemical Ilnd of mechanical menns has been restudied by War. burton (1058), Cobh (1069) and Rtitzlcr llnd Rieger (1973). It is now accepted that the excavation ill performed by cellular etching of the substratum, which results in freeing of characteristically shaped calcium carbonates chips Wig. I b and c). These chips are expelled through the oscula and are carried away by water currents. Although the nature of the etching agent (possibly carbonic anhydrase) is still under investigation (Hatch, 1972). it has boon calculated that only 2-3% of the eroded subslratum is removed in dissolved form (Rtitzler and Rieger, 1973). Otter (1973) and Ginsburg (1957) pointed to tho fact that sponge burrowing plays an important part in weakening of t.he coral reef framework, thus accelerating erosion by wave act·ion in shallow water. Goreau and Itartrnan (1963) demon­ strated much more obvious destructive effect-s caused by sponge burrowing on the fore-reef slope in Jamaica, below 30 m. They also suggested t.hat these sponges contribute to t.he large mud fraction of ee<liment8 produced in roof communities. This latter assumption was further supported by Neumllnn (1965, 1966) at Bermuda lind by S. V. Smith (personnl communication, 1972) at .Fanning Island. Fiitterer (1974) determined and counted clionid-produced partieles by scanning electron microscopy. He calculated that sponge.generated fine sediment& can amount to 2-3% (Northern Adriatic Sea, Persian Gulf) or even to 30% (Fanning Island) of the total sediment. Rates of biocrosioD caused by Cliona tampa were determined by Neumann (1966) by measuring weight loss of limestone blocks which had been attached to sponge bearing material for up to 100 days. From this data he calculated that. CliQna infested substrata can lose 8S much as 1.4 cm per year, or 6-7 kg per m2 per 100 days, producing nearly 6 kg fine sediments per m2 per 100 days (assuming 10% removal in solution, as proposed by Warburton, 1958). During a recent study 011 the burrowing meehallism of Climl.a lampa (Riitzlcr and Rieger, 1973), I initiated similar experiments to obtain new growth of this sponge ill blocks of Iceland spar. I calculated comparably high penetration rates but was puzzled by the fact that labeled and mCll.8ured "old" CliQna~inrested substrata (mollusk shells, small coral heads and limestone rock) had not shown significant shape or size changes during 16 months of in situ observation. My conclusion, that a stimulation effect would cause exceptionally high rates during the invasion of new substrata had also been suggested by Neumann (1966). Measurement& of long-term excavation rates are, therefore, important for realistic estimates of sponge participation in eros.ion and sediment production. For this purpose I left calcite blocks. attached to Cliona tampa in Bermuda. for up to 10 months during 1972-1!)74. A 16-month experiment with particularly large blocks to offer ample new substratum was unfortunately lost by storm The Role of Burrowing Spongea in Biool'06io/l 2{)5 or vandalism. A similar 12.month experiment, however, conducted with conch shells attached to Cliana aprim at the Belize (British Honduras) barrier reef (1972/1973) waa succcssfui. To monit-or weight loss of materials already infested by sponges, weighed and labeled bivalve shells containing Cliona lampa were returned to their previous habitats for 7 months (1973). Short term burrowing activity patterns and possible influence of temperature and water movement were also investigated. For this part of the study, a dense population of Cliona lampa growing on isolated concrete piJIars in the center of Ferry Reach Entrance (North Shore, Bermuda) was used. 'fhe sponges infest extensive accumulations of the bivalve Chama 7/w.«rophylla (Fig. I a) which fouls the pillars. The tides cause a directional flow of water in the narrow channel. Thus, the characteristically shaped chips produced by the sponges could be trapped in sediment jars during periods of observation in January and August, 1973. They were measured and counted in comparison with controls outside the influence of the OUona population. 'fo evaluate the ecological consequences of the burrowing activity, it is necessary to rclat-e the rates of erosion and sediment production to sponge abun. dance in a given area. For this purpose, the relative abundance of excavating sponge species hitherto known from Bermuda (Riitzler, 1974) was determined by a series of representat.ive t.ransect coullta. ~lateri8ls and Uethods .For the excavlltion experimenta in Bermuda pre.weighed Iceland spar blooks were attached to Cliona lampa Laubenfels (forma lampa) under water (1-3 m) using plastic coated wire. The procedures follo,,'ed those outlined by Rutzler and Rieger (1973). The blocks selected had a size clO8C to 3 X2 X 1.5 cm. In the reef crest area of ClIrric Bow Cay, Belize, slabs chiseled from fresh shells of Slrombus gifIM UnMeus were weighed and wired to Cliona. aprial Psng (forma aprka) which burrowed the coral Aeropora palmala (Lamarck) in about I-2m depth. The conch slabs ranged in size 6x4xO.5cm to 12xlOx1.8cm, After the experiments the area of initial penetration by the sponge (= contract area) and the area of total sponge coverage (ne"' gro",th) were measured using plsnimetry on pllper tracings. Then the sponge material Willi removed by soaking in commercial lIOdium hypochloride. ClIlcified fouling organisms were removed by wreful scrnping. After rinsing and removing spiculell and debris with s jet of water, the blooks were dried and weighed. To record burrowing llctivity patterns 8 sediment lraps were placed between 4- large concrete pillai'll in Ferry !'leach Entrnnoo (Bermuda). The pillai'll are fouled by a dell&6 population of the bivalve Clwmo. m~roph.ylla Gmelin, which is infested by CliQlI(J lompa. The CHOlia population pl'el>6nt above and surrounding the traps was calculated to cover 62 m!. within a 5 m radius. A fair portion of cllipll produced by these spongea could, l·herefore, be trnpped using 500 ml gla8& jnl'll with 60 mm diameter openings.
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